功能多样性及其研究方法

功能多样性是联系生物多样性和生态系统功能的关键性因素。植物群落内功能多样性的量化描述对于研究生物多样性响应环境变化及其对生物多样性-生态系统功能关系的影响至关重要。由于功能多样性的概念含混不清,使得对其范围的确定、定量描述及其对生态系统功能的作用的评价存在较大的难度。在查阅大量文献的基础上,以植物群落为例,介绍了功能多样性的定义、计算方法及研究中应注意的问题。在植物群落功能多样性指数的选择和测定过程中,对获取相关物种必要的功能特征信息、确定其权重、确定不同物种在群落中的相对丰富度、利用所得的功能多样性指数解释和预测生态系统过程的变化等问题,进行了研究。     

Drivers of productivity and its temporal stability in a tropical tree diversity experiment

There is increasing evidence that mixed‐species forests can provide multiple ecosystem services at a higher level than their monospecific counterparts. However, most studies concerning tree diversity and ecosystem functioning relationships use data from forest inventories (under noncontrolled conditions) or from very young plantation experiments. Here, we investigated temporal dynamics of diversity–productivity relationships and diversity–stability relationships in the oldest tropical tree diversity experiment. Sardinilla was established in Panama in 2001, with 22 plots that form a gradient in native tree species richness of one‐, two‐, three‐ and five‐species communities. Using annual data describing tree diameters and heights, we calculated basal area increment as the proxy of tree productivity. We combined tree neighbourhood‐ and community‐level analyses and tested the effects of both species diversity and structural diversity on productivity and its temporal stability. General patterns were consistent across both scales indicating that tree–tree interactions in neighbourhoods drive observed diversity effects. From 2006 to 2016, mean overyielding (higher productivity in mixtures than in monocultures) was 25%–30% in two‐ and three‐species mixtures and 50% in five‐species stands. Tree neighbourhood diversity enhanced community productivity but the effect of species diversity was stronger and increased over time, whereas the effect of structural diversity declined. Temporal stability of community productivity increased with species diversity via two principle mechanisms: asynchronous responses of species to environmental variability and overyielding. Overyielding in mixtures was highest during a strong El Niño‐related drought. Overall, positive diversity–productivity and diversity–stability relationships predominated, with the highest productivity and stability at the highest levels of diversity. These results provide new insights into mixing effects in diverse, tropical plantations and highlight the importance of analyses of temporal dynamics for our understanding of the complex relationships between diversity, productivity and stability. Under climate change, mixed‐species forests may provide both high levels and high stability of production.     

Functional diversity of crustacean zooplankton communities: towards a trait-based classification

1. While studies of phytoplankton and terrestrial plant communities have increasingly emphasised the use of functional traits in ecological research, few have yet to apply this approach to zooplankton communities.
2. This study reviews laboratory and observational studies on zooplankton feeding and life history and provides a series of functional trait tables for the North American freshwater zooplankton. Qualitative and quantitative trait tables highlight areas where data were more scarce and point to which types of studies could fill in gaps in our knowledge of zooplankton niches.
3. Data were most complete for the Cladocera across most traits, while feeding information for cyclopoids was most sparse. Qualitative data that distinguished congeneric species were lacking for most groups.
4. A regional community dendrogram for common north-eastern North American zooplankton species was generated and shows that taxonomic differences between species do not capture fully functional differences based on the traits of body length, habitat, trophic group and feeding type.
5. The data collected here, combined with readily measurable species attributes, can be used to generate a multivariate measure of the functional niche of each species found in a community. Armed with this information, functional relationships that are useful for ecological studies of lake ecosystems can be more easily conducted.     

From competition to facilitation: how tree species respond to neighbourhood diversity

Studies on tree communities have demonstrated that species diversity can enhance forest productivity, but the driving mechanisms at the local neighbourhood level remain poorly understood. Here, we use data from a large‐scale biodiversity experiment with 24 subtropical tree species to show that neighbourhood tree species richness generally promotes individual tree productivity. We found that the underlying mechanisms depend on a focal tree's functional traits: For species with a conservative resource‐use strategy diversity effects were brought about by facilitation, and for species with acquisitive traits by competitive reduction. Moreover, positive diversity effects were strongest under low competition intensity (quantified as the total basal area of neighbours) for acquisitive species, and under high competition intensity for conservative species. Our findings demonstrate that net biodiversity effects in tree communities can vary over small spatial scales, emphasising the need to consider variation in local neighbourhood interactions to better understand effects at the community level.     

Mixing effects on litter decomposition rates in a young tree diversity experiment

Litter decomposition is an essential process for biogeochemical cycling and for the formation of new soil organic matter. Mixing litter from different tree species has been reported to increase litter decomposition rates through synergistic effects. We assessed the decomposition rates of leaf litter from five tree species in a recently established tree diversity experiment on a post-agriculture site in Belgium. We used 20 different leaf litter compositions with diversity levels ranging from 1 up to 4 species. Litter mass loss in litterbags was assessed 10, 20, 25, 35, and 60 weeks after installation in the field. We found that litter decomposition rates were higher for high-quality litters, i.e., with high nitrogen content and low lignin content. The decomposition rates of mixed litter were more affected by the identity of the litter species within the mixture than by the diversity of the litter per se, but the variability in litter decomposition rates decreased as the litter diversity increased. Among the 15 different mixed litter compositions in our study, only three litter combinations showed synergistic effects. Our study suggests that admixing tree species with high-quality litter in post-agricultural plantations helps in increasing the mixture's early-stage litter decomposition rate.     

Response diversity determines the resilience of ecosystems to environmental change

A growing body of evidence highlights the importance of biodiversity for ecosystem stability and the maintenance of optimal ecosystem functionality. Conservation measures are thus essential to safeguard the ecosystem services that biodiversity provides and human society needs. Current anthropogenic threats may lead to detrimental (and perhaps irreversible) ecosystem degradation, providing strong motivation to evaluate the response of ecological communities to various anthropogenic pressures. In particular, ecosystem functions that sustain key ecosystem services should be identified and prioritized for conservation action. Traditional diversity measures (e.g. ‘species richness’) may not adequately capture the aspects of biodiversity most relevant to ecosystem stability and functionality, but several new concepts may be more appropriate. These include ‘response diversity’, describing the variation of responses to environmental change among species of a particular community. Response diversity may also be a key determinant of ecosystem resilience in the face of anthropogenic pressures and environmental uncertainty. However, current understanding of response diversity is poor, and we see an urgent need to disentangle the conceptual strands that pervade studies of the relationship between biodiversity and ecosystem functioning. Our review clarifies the links between response diversity and the maintenance of ecosystem functionality by focusing on the insurance hypothesis of biodiversity and the concept of functional redundancy. We provide a conceptual model to describe how loss of response diversity may cause ecosystem degradation through decreased ecosystem resilience. We explicitly explain how response diversity contributes to functional compensation and to spatio‐temporal complementarity among species, leading to long‐term maintenance of ecosystem multifunctionality. Recent quantitative studies suggest that traditional diversity measures may often be uncoupled from measures (such as response diversity) that may be more effective proxies for ecosystem stability and resilience. Certain conclusions and recommendations of earlier studies using these traditional measures as indicators of ecosystem resilience thus may be suspect. We believe that functional ecology perspectives incorporating the effects and responses of diversity are essential for development of management strategies to safeguard (and restore) optimal ecosystem functionality (especially multifunctionality). Our review highlights these issues and we envision our work generating debate around the relationship between biodiversity and ecosystem functionality, and leading to improved conservation priorities and biodiversity management practices that maximize ecosystem resilience in the face of uncertain environmental change.     

Herbivore and pathogen effects on tree growth are additive,but mediated by tree diversity and plant traits

Herbivores and fungal pathogens are key drivers of plant community composition and functioning. The effects of herbivores and pathogens are mediated by the diversity and functional characteristics of their host plants. However, the combined effects of herbivory and pathogen damage, and their consequences for plant performance, have not yet been addressed in the context of biodiversity–ecosystem functioning research. We analyzed the relationships between herbivory, fungal pathogen damage and their effects on tree growth in a large‐scale forest‐biodiversity experiment. Moreover, we tested whether variation in leaf trait and climatic niche characteristics among tree species influenced these relationships. We found significant positive effects of herbivory on pathogen damage, and vice versa. These effects were attenuated by tree species richness—because herbivory increased and pathogen damage decreased with increasing richness—and were most pronounced for species with soft leaves and narrow climatic niches. However, herbivory and pathogens had contrasting, independent effects on tree growth, with pathogens decreasing and herbivory increasing growth. The positive herbivory effects indicate that trees might be able to (over‐)compensate for local damage at the level of the whole tree. Nevertheless, we found a dependence of these effects on richness, leaf traits and climatic niche characteristics of the tree species. This could mean that the ability for compensation is influenced by both biodiversity loss and tree species identity—including effects of larger‐scale climatic adaptations that have been rarely considered in this context. Our results suggest that herbivory and pathogens have additive but contrasting effects on tree growth. Considering effects of both herbivory and pathogens may thus help to better understand the net effects of damage on tree performance in communities differing in diversity. Moreover, our study shows how species richness and species characteristics (leaf traits and climatic niches) can modify tree growth responses to leaf damage under real‐world conditions.     

Sick plants in grassland communities: a growth‐defense trade‐off is the main driver of fungal pathogen abundance

Aboveground fungal pathogens can substantially reduce biomass production in grasslands. However, we lack a mechanistic understanding of the drivers of fungal pathogen infection and impact. Using a grassland global change and biodiversity experiment we show that the trade‐off between plant growth and defense is the main determinant of infection incidence. In contrast, nitrogen addition only indirectly increased incidence via shifting plant communities towards faster growing species. Plant diversity did not decrease incidence, likely because spillover of generalist pathogens or dominance of susceptible plants counteracted negative diversity effects. A fungicide treatment increased plant biomass production and high levels of infection incidence were associated with reduced biomass. However, pathogen impact was context dependent and infection incidence reduced biomass more strongly in diverse communities. Our results show that a growth‐defense trade‐off is the key driver of pathogen incidence, but pathogen impact is determined by several mechanisms and may depend on pathogen community composition.     

Ant diversity as a direct and indirect driver of pselaphine rove beetle (Coleoptera: Staphylinidae) functional diversity in tropical rainforests,Sabah, Malaysian Borneo

Pselaphinae is a species‐rich beetle subfamily found globally, with many exhibiting myrmecophily—a symbiotic association with ants. Pselaphine–ant associations vary from facultative to obligate, but direct behavioral observations still remain scarce. Pselaphines are speciose and ecologically abundant within tropical leaf litter invertebrate communities where ants dominate, implying a potentially important ecological role that may be affected by habitat disturbances that impact ants. In this study, we measured and analyzed putative functional traits of leaf litter pselaphines associated with myrmecophily through morphometric analysis. We calculated “myrmecophile functional diversity” of pselaphines at different sites and examined this measure's relationship with ant abundance, in both old growth and logged rainforest sites in Sabah, Borneo. We show that myrmecophile functional diversity of pselaphine beetles increases as ant abundance increases. Old growth rainforest sites support a high abundance of ants, which is associated with a high abundance of probable myrmecophilous pselaphines. These results suggest a potential link between adult morphological characters and the functional role these beetles play in rainforest litter as ecological interaction partners with ants.     

Partitioning the biodiversity effects on productivity into density and size components

Plant density and size — two factors that represent plant survival and growth — are key determinants of yield but have rarely been analysed explicitly in the context of biodiversity–productivity relationships. Here, we derive equations to partition the net, complementarity and selection effects of biodiversity into additive components that reflect diversity-induced changes in plant density and size. Applications of the new method to empirical datasets reveal contrasting ways in which plant density and size regulate yield in species mixtures. In an annual plant diversity experiment, overyielding is largely explained by selection effects associated with increased size of highly productive plant species. In a tree diversity experiment, the cause of overyielding shifts from enhanced growth in tree size to reduced mortality by complementary use of canopy space during stand development. These results highlight the capability of the new method to resolve crucial, yet understudied, demographic links between biodiversity and productivity.     

Hollow oaks and beetle functional diversity: Significance of surroundings extends beyond taxonomy

Veteran hollow oaks (Quercus spp.) are keystone structures hosting high insect diversity but are declining in numbers due to intensification of land use and the abandonment of traditional management. The loss of this vital habitat is resulting in a reduction of biodiversity, and this likely has consequences for ecosystem functioning, especially if functional diversity is reduced. A considerable amount of research has been done on predictors of beetle taxonomic diversity in veteran oaks, but predictors of functional diversity have remained largely unexplored. The aim of this study was to establish whether the features and surroundings of veteran oaks are related to functional diversity within three functional groups of beetles (decomposers, predators, and flower visitors) and determine whether species richness and functional diversity within the groups are dependent on the same predictors. Sampling was carried out intermittently between 2004 and 2011 on 61 veteran oaks in Southern Norway. Of the 876 beetle species that were collected, 359 were determined to be decomposers, 284 were predators, and 85 were flower visitors. Species richness and functional diversity in all groups were consistently higher in traps mounted on veteran oaks in forests than in open landscapes. However, additional predictors differed between groups, and for species richness and functional diversity. Decomposer species richness responded to tree vitality, while functional diversity responded to habitat connectivity, predator species richness responded to regrowth of shrubs while functional diversity responded to tree circumference, and flower visitor richness and functional diversity did not respond to any additional predictors. Previous studies have found that the features and surroundings of veteran oaks are important for conservation of taxonomic diversity, and the results from this study indicate that they are also important for functional diversity within multiple functional groups.     

Traits of dominant plant species drive normalized difference vegetation index in grasslands globally

Aim

Theoretical, experimental and observational studies have shown that biodiversity–ecosystem functioning (BEF) relationships are influenced by functional community structure through two mutually non-exclusive mechanisms: (1) the dominance effect (which relates to the traits of the dominant species); and (2) the niche partitioning effect [which relates to functional diversity (FD)]. Although both mechanisms have been studied in plant communities and experiments at small spatial extents, it remains unclear whether evidence from small-extent case studies translates into a generalizable macroecological pattern. Here, we evaluate dominance and niche partitioning effects simultaneously in grassland systems world-wide.

Location

Two thousand nine hundred and forty-one grassland plots globally.

Time period

2000–2014.

Major taxa studied

Vascular plants.

Methods

We obtained plot-based data on functional community structure from the global vegetation plot database “sPlot”, which combines species composition with plant trait data from the “TRY” database. We used data on the community-weighted mean (CWM) and FD for 18 ecologically relevant plant traits. As an indicator of primary productivity, we extracted the satellite-derived normalized difference vegetation index (NDVI) from MODIS. Using generalized additive models and deviation partitioning, we estimated the contributions of trait CWM and FD to the variation in annual maximum NDVI, while controlling for climatic variables and spatial structure.

Results

Grassland communities dominated by relatively tall species with acquisitive traits had higher NDVI values, suggesting the prevalence of dominance effects for BEF relationships. We found no support for niche partitioning for the functional traits analysed, because NDVI remained unaffected by FD. Most of the predictive power of traits was shared by climatic predictors and spatial coordinates. This highlights the importance of community assembly processes for BEF relationships in natural communities.

Main conclusions

Our analysis provides empirical evidence that plant functional community structure and global patterns in primary productivity are linked through the resource economics and size traits of the dominant species. This is an important test of the hypotheses underlying BEF relationships at the global scale.     

Early subtropical forest growth is driven by community mean trait values and functional diversity rather than the abiotic environment

While functional diversity (FD) has been shown to be positively related to a number of ecosystem functions including biomass production, it may have a much less pronounced effect than that of environmental factors or species‐specific properties. Leaf and wood traits can be considered particularly relevant to tree growth, as they reflect a trade‐off between resources invested into growth and persistence. Our study focussed on the degree to which early forest growth was driven by FD, the environment (11 variables characterizing abiotic habitat conditions), and community‐weighted mean (CWM) values of species traits in the context of a large‐scale tree diversity experiment (BEF‐China). Growth rates of trees with respect to crown diameter were aggregated across 231 plots (hosting between one and 23 tree species) and related to environmental variables, FD, and CWM, the latter two of which were based on 41 plant functional traits. The effects of each of the three predictor groups were analyzed separately by mixed model optimization and jointly by variance partitioning. Numerous single traits predicted plot‐level tree growth, both in the models based on CWMs and FD, but none of the environmental variables was able to predict tree growth. In the best models, environment and FD explained only 4 and 31% of variation in crown growth rates, respectively, while CWM trait values explained 42%. In total, the best models accounted for 51% of crown growth. The marginal role of the selected environmental variables was unexpected, given the high topographic heterogeneity and large size of the experiment, as was the significant impact of FD, demonstrating that positive diversity effects already occur during the early stages in tree plantations.     

Experimental evidence for the effects of dung beetle functional group richness and composition on ecosystem function in a tropical forest

Much of the literature on the relationship between species richness or functional group richness and measures of ecosystem function focuses on a restricted set of ecosystem function measures and taxonomic groups. Few such studies have been carried out under realistic levels of diversity in the field, particularly in high diversity ecosystems such as tropical forests. We used exclusion experiments to study the effects of dung beetle functional group richness and composition on two interlinked and functionally important ecological processes, dung removal and secondary seed dispersal, in evergreen tropical forest in Sabah, Malaysian Borneo. Overall, both dung and seed removal increased with dung beetle functional group richness. However, levels of ecosystem functioning were idiosyncratic depending on the identity of the functional groups present, indicating an important role for functional group composition. There was no evidence for interference or competition among functional groups. We found strong evidence for overyielding and transgressive overyielding, suggesting complementarity or facilitation among functional groups. Not all mixtures showed transgressive overyielding, so that complementarity was restricted to particular functional group combinations. Beetles in a single functional group (large nocturnal tunnellers) had a disproportionate influence on measures of ecosystem function: in their absence dung removal is reduced by approximately 75%. However, a full complement of functional groups is required to maximize ecosystem functioning. This study highlights the importance of both functional group identity and species composition in determining the ecosystem consequences of extinctions or altered patterns in the relative abundance of species.     

Forest gene diversity is correlated with the composition and function of soil microbial communities

The growing field of community and ecosystem genetics indicates that plant genotype and genotypic variation are important for structuring communities and ecosystem processes. Little is known, however, regarding the effects of stand gene diversity on soil communities and processes under field conditions. Utilizing natural genetic variation occurring in Populus spp. hybrid zones, we tested the hypothesis that stand gene diversity structures soil microbial communities and influences soil nutrient pools. We found significant unimodal patterns relating gene diversity to soil microbial community composition, microbial exoenzyme activity of a carbon-acquiring enzyme, and availability of soil nitrogen. Multivariate analyses indicate that this pattern is due to the correlation between gene diversity, plant secondary chemistry, and the composition of the microbial community that impacts the availability of soil nitrogen. Together, these data from a natural system indicate that stand gene diversity may affect soil microbial communities and soil processes in ways similar to species diversity (i.e., unimodal patterns). Our results further demonstrate that the effects of plant genetic diversity on other organisms may be mediated by plant functional trait variation.